Laura Yahdjian

Water pulses and biogeochemical cycles in arid and semiarid ecosystems

The episodic nature of water availability in arid and semiarid ecosystems has significant consequences on belowground carbon and nutrient cycling. Pulsed water events directly control belowground processes through soil wet-dry cycles. Rapid soil microbial response to incident moisture availability often results in almost instantaneous C and N mineralization, followed by shifts in C/N of microbially available substrate, and an offset in the balance between nutrient immobilization and mineralization. Nitrogen inputs from biological soil crusts are also highly sensitive to pulsed rain events, and nitrogen losses, particularly gaseous losses due to denitrification and nitrate leaching, are tightly linked to pulses of water availability. The magnitude of the effect of water pulses on carbon and nutrient pools, however, depends on the distribution of resource availability and soil organisms, both of which are strongly affected by the spatial and temporal heterogeneity of vegetation cover, topographic position and soil texture. The ‘inverse texture hypothesis’ for net primary production in water-limited ecosystems suggests that coarse-textured soils have higher NPP than fine-textured soils in very arid zones due to reduced evaporative losses, while NPP is greater in fine-textured soils in higher rainfall ecosystems due to increased water-holding capacity. With respect to belowground processes, fine-textured soils tend to have higher water-holding capacity and labile C and N pools than coarse-textured soils, and often show a much greater flush of N mineralization. The result of the interaction of texture and pulsed rainfall events suggests a corollary hypothesis for nutrient turnover in arid and semiarid ecosystems with a linear increase of N mineralization in coarse-textured soils, but a saturating response for fine-textured soils due to the importance of soil C and N pools. Seasonal distribution of water pulses can lead to the accumulation of mineral N in the dry season, decoupling resource supply and microbial and plant demand, and resulting in increased losses via other pathways and reduction in overall soil nutrient pools. The asynchrony of resource availability, particularly nitrogen versus water due to pulsed water events, may be central to understanding the consequences for ecosystem nutrient retention and long-term effects on carbon and nutrient pools. Finally, global change effects due to changes in the nature and size of pulsed water events and increased asynchrony of water availability and growing season will likely have impacts on biogeochemical cycling in water-limited ecosystems.

A rainout shelter design for intercepting different amounts of rainfall

Abstract. Field manipulative experiments represent a straightforward way to explore temporal relationships between annual precipitation and productivity. Water exclusion usually involves the use of rainout shelters, which are in general formed by a complete roof that intercepts 100% of the rainfall and require complicated mechanisms to move the shelter into place. The rainout-shelter design described here is a fixed-location shelter with a roof consisting of bands of transparent acrylic that blocks different amounts of rainfall while minimally affecting other environmental variables. We constructed thirty 3.76-m2 shelters in an arid steppe near Río Mayo, Argentina (at 45°41′S, 70°16′W), to impose 30%, 55%, and 80% of rainfall interception. We tested the effectiveness of the design by collecting all the intercepted water in storage tanks and we evaluated changes in soil water content with the time domain reflectometry technique. We also measured soil water content in regular grids to assess the magnitude of the edge effect. We analysed the microclimate impact of the shelters by measuring photosynthetically active radiation and air and soil temperature inside and outside shelters. We did not detect significant differences between the observed and the expected rainfall interception for the 30% and 55% interception treatments but the 80% shelters intercepted 71% of incoming rainfall, which was significantly (P<0.05) lower than the expected value. Soil water content was significantly (P<0.05) higher in the control plots than in the plots with rainout shelter at all dates, except in January (summer). Radiation was not affected by the 30% interception treatment, but the roof with the largest number of acrylics bands (80% interception treatment) reduced incident radiation throughout the day by 10%. Air and soil temperatures were lower under than outside the shelters during the period of highest radiation but the opposite occurred with low radiation but with smaller differences. The two characteristics of the shelter, fixed design and low cost, allow for proper replication in space, which is required in ecological field experiments.

VEGETATION STRUCTURE CONSTRAINS PRIMARY PRODUCTION RESPONSE TO WATER AVAILABILITY IN THE PATAGONIAN STEPPE

Grassland aboveground net primary production (ANPP) increases linearly with precipitation in space and time, but temporal models relating time series of ANPP and annual precipitation for single sites show lower slopes and regression coefficients than are shown by spatial models. The analysis of several ANPP time series showed lags in the ecosystem response to increased water availability, which may explain the difference between spatial and temporal models. The lags may result from constraints that ecosystems experience after drought. Our objective was to explore the structural constraints of the ANPP response to rainfall variability in a semiarid ecosystem, the Patagonian steppe, in southern Argentina. We designed a 3-yr rainfall manipulation experiment where we decreased water input with rainout shelters during two consecutive years, which included three levels of rainfall interception (30%, 55%, and 80%) and a control. In the third year, we irrigated one-half of the plots of each rainfall-interception treatment. We evaluated the immediate effects of drought on current-year ANPP and the effects of previous-year drought on vegetation recovery after water supplementation. ANPP (g x m(-2) x yr(-1)) was linearly related to annual precipitation input (APPT; mm/yr) along the experimental precipitation gradient (ANPP = 0.13 x APPT + 58.3; r2 = 0.34, P < 0.01), and this relationship was mostly accounted for by changes in the ANPP of grasses. Plant density (D; no. individuals/mm2) was related to the precipitation received during the drought period (D = 0.11 x APPT + 18; r2 = 0.39, P < 0.05). The recovery of plants after irrigation was lower for those plots that had experienced experimental drought the previous years relative to controls, and the lags were proportional to the intensity of drought. Therefore, our results suggest that the density of plants may constrain the recovery of vegetation after drought, and these constraints may determine lags that limit the capacity of the ecosystem to take advantage of wet years after dry years.

Differential Controls of Water Input on Litter Decomposition and Nitrogen Dynamics in the Patagonian Steppe

Studies of the effects of precipitation on litter decomposition and nitrogen mineralization in arid and semiarid environments have demonstrated contradictory results. We conducted a manipulative experiment with rainout shelters in the semiarid Patagonian steppe, aimed at assessing the direct effects of water availability on litter decomposition and net nitrogen mineralization while isolating the indirect effects. We created four levels of precipitation input: control and three levels (30, 55 and 80%) of precipitation interception and we examined litter decomposition and nutrient release of a dominant grass species, Stipa speciosa, inorganic soil nitrogen, and in situ net nitrogen mineralization over two consecutive years. Litter decomposition rates (k, year−1) varied significantly (P < 0.001) among precipitation interception treatments and were positively correlated with incoming annual precipitation (APPT, mm/year) (k = 0.0007 × APPT + 0.137). In contrast, net N mineralization was not correlated with incoming precipitation. Soil NO3− significantly decreased with increasing precipitation input, whereas soil NH4+ concentration did not differ among precipitation interception treatments. Controls of water input on litter decomposition appear to be different from controls on N mineralization in the semiarid Patagonian steppe. We suggest that although water availability affects both the mineralization of C and N, it differentially affects the movement and fate of the inorganic products. A consequence of the accumulation of inorganic N during dry episodes is that periods of maximum water and soil nutrient availability may occur at different times. This asynchrony in the availability of N and water in the soil may explain the observed lags in the response of primary production to increases in water availability.

Asymmetric responses of primary productivity to precipitation extremes: A synthesis of grassland precipitation manipulation experiments

Climatic changes are altering Earths hydrological cycle, resulting in altered precipitation amounts, increased interannual variability of precipitation, and more frequent extreme precipitation events. These trends will likely continue into the future, having substantial impacts on net primary productivity (NPP) and associated ecosystem services such as food production and carbon sequestration. Frequently, experimental manipulations of precipitation have linked altered precipitation regimes to changes in NPP. Yet, findings have been diverse and substantial uncertainty still surrounds generalities describing patterns of ecosystem sensitivity to altered precipitation. Additionally, we do not know whether previously observed correlations between NPP and precipitation remain accurate when precipitation changes become extreme. We synthesized results from 83 case studies of experimental precipitation manipulations in grasslands worldwide. We used meta-analytical techniques to search for generalities and asymmetries of aboveground NPP (ANPP) and belowground NPP (BNPP) responses to both the direction and magnitude of precipitation change. Sensitivity (i.e., productivity response standardized by the amount of precipitation change) of BNPP was similar under precipitation additions and reductions, but ANPP was more sensitive to precipitation additions than reductions; this was especially evident in drier ecosystems. Additionally, overall relationships between the magnitude of productivity responses and the magnitude of precipitation change were saturating in form. The saturating form of this relationship was likely driven by ANPP responses to very extreme precipitation increases, although there were limited studies imposing extreme precipitation change, and there was considerable variation among experiments. This highlights the importance of incorporating gradients of manipulations, ranging from extreme drought to extreme precipitation increases into future climate change experiments. Additionally, policy and land management decisions related to global change scenarios should consider how ANPP and BNPP responses may differ, and that ecosystem responses to extreme events might not be predicted from relationships found under moderate environmental changes.

Preference for different inorganic nitrogen forms among plant functional types and species of the Patagonian steppe

We have explored species–specific preferences for nitrate (NO3−) and ammonium (NH4+) as an alternative niche separation in ecosystems where nitrogen (N) is present mostly in inorganic forms. The Patagonian steppe is dominated by shrubs and grasses. Shrubs absorb water and nutrients from deep soil layers, which are poor in N, while grasses have the opposite pattern, absorbing most of their water and nutrients from the upper layers of the soil. We hypothesized that the preferences of shrub and grass for inorganic N forms are different and that the rate of potential N uptake is greater in shrubs than in grasses. To test this hypothesis, we grew individuals of six dominant species in solutions of different NH4+:NO3− concentration ratios. Nitrate uptake was found to be higher in shrubs, while ammonium uptake was similar between plant functional types. The NH4+:NO3− uptake ratio was significantly lower for shrubs than grasses. Shrubs, which under field conditions have deeper rooting systems than grasses, showed a higher N absorption capacity than grasses and a preference for the more mobile N form, nitrate. Grasses, which had lower N uptake rates than shrubs, preferred ammonium over nitrate. These complementary patterns between grasses and shrubs suggest a more thorough exploitation of resources by diverse ecosystems than those dominated by just one functional type. The loss of one plant functional group or a significant change in its abundance would therefore represent a reduction in resource use efficiency and ecosystem functioning.

Climate modifies response of non-native and native species richness to nutrient enrichment

Ecosystem eutrophication often increases domination by non-natives and causes displacement of native taxa. However, variation in environmental conditions may affect the outcome of interactions between native and non-native taxa in environments where nutrient supply is elevated. We examined the interactive effects of eutrophication, climate variability and climate average conditions on the success of native and non-native plant species using experimental nutrient manipulations replicated at 32 grassland sites on four continents. We hypothesized that effects of nutrient addition would be greatest where climate was stable and benign, owing to reduced niche partitioning. We found that the abundance of non-native species increased with nutrient addition independent of climate; however, nutrient addition increased non-native species richness and decreased native species richness, with these effects dampened in warmer or wetter sites. Eutrophication also altered the time scale in which grassland invasion responded to climate, decreasing the importance of long-term climate and increasing that of annual climate. Thus, climatic conditions mediate the responses of native and non-native flora to nutrient enrichment. Our results suggest that the negative effect of nutrient addition on native abundance is decoupled from its effect on richness, and reduces the time scale of the links between climate and compositional change.

Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands—those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)—had higher levels of multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities.Analysis of 65 grasslands worldwide from the Nutrient Network experiment reveals that plant communities with higher α- and β-diversity have higher levels of ecosystem multifunctionality, and that this effect is amplified across scales.

Rangeland Ecosystem Services: Nature’s Supply and Humans’ Demand

Ecosystem services are the benefits that society receives from nature, including the regulation of climate, the pollination of crops, the provisioning of intellectual inspiration and recreational environment, as well as many essential goods such as food, fiber, and wood. Rangeland ecosystem services are often valued differently by different stakeholders interested in livestock production, water quality and quantity, biodiversity conservation, or carbon sequestration. The supply of ecosystem services depends on biophysical conditions and land-use history, and their availability is assessed using surveys of soils, plants, and animals. The demand for ecosystem services depends on educational level, income, and location of residence of social beneficiaries. The demand can be assessed through stakeholder interviews, questionnaires, and surveys. Rangeland management affects the supply of different ecosystem services by producing interactions among them. Trade-offs result when an increase in one service is associated with a decline in another, and win–win situations occur when an increase in one service is associated with an increase in other services. This chapter provides a conceptual framework in which range management decisions are seen as a challenge of reconciling supply and demand of ecosystem services.

Global-change drivers of ecosystem functioning modulated by natural variability and saturating responses.

Humans are altering global environment at an unprecedented rate through changes in biodiversity, climate, nitrogen cycle, and land use. To address their effects on ecosystem functioning, experiments most frequently explore one driver at a time and control as many confounding factors as possible. Yet, which driver exerts the largest influence on ecosystem functioning and whether their relative importance changes among systems remain unclear. We analyzed experiments in the Patagonian steppe that evaluated the aboveground net primary production (ANPP) response to manipulated gradients of species richness, precipitation, temperature, nitrogen fertilization (N), and grazing intensity. We compared the effect on ANPP relative to ambient conditions considering intensity and direction of manipulations for each driver. The ranking of responses to drivers with comparable manipulation intensity was as follows: biodiversity>grazing>precipitation>N. For a similar intensity of manipulation, the effect of biodiversity loss was 4.0, 3.6, and 1.5, times larger than N deposition, decreased precipitation, and increased grazing intensity. We interpreted our results considering two hypotheses. First, the response of ANPP to changes in precipitation and biodiversity is saturating, so we expected larger effects when the driver was reduced, relative to ambient conditions, than when it was increased. Experimental manipulations that reduced ambient levels had larger effects than those that increased them. Second, the sensitivity of ANPP to each driver is inversely related to the natural variability of the driver. In Patagonia, the ranking of natural variability of drivers is as follows: precipitation>grazing>temperature>biodiversity>N. So, in general, the ecosystem was most sensitive to drivers that varied the least. Comparable results from Cedar Creek (MN) support both hypotheses and suggest that sensitivity to drivers varies among ecosystem types. Given the importance of understanding ecosystem sensitivity to predict global-change impacts, it is necessary to design new experiments located in regions with contrasting natural variability and that include the full range of drivers.